Variability of venom components in immune suppressive parasitoid wasps: from a phylogenetic to a population approach.

Endoparasitoid wasps develop at the expense of other insects, leading to their death. Eggs deposited inside the host body induce an immune response, which results in the formation of a melanized cellular capsule around the egg. To evade or counteract this response, endoparasitoids have evolved different strategies, the most often reported being injection into the host of immunosuppressive factors, notably venom proteins, along with the egg. The analysis of venom components has been performed independently in species of different taxa, but the present picture is far from complete. Intriguingly, the question of the level of venom variability inside species has been neglected, although it may partly determine the potential for parasitoid adaptation. Here, we present a short review of our present knowledge of venom components in endoparasitoids, as well as of the only well-known example of intraspecific variability in a venom immune suppressive protein being responsible for variation in parasitoid virulence. We then present data evidencing inter-individual variation of venom protein profiles, using a gel electrophoresis approach, both in laboratory strains and field populations of a figitid and a braconid species. Whether occurrence of such variability may permit a selection of parasitoid venom components driven by the host remains to be tested, notably in the context of the production and use of biological control auxiliaries.

Geographic variations of life history traits and potential trade-offs in different populations of the parasitoid Leptopilina heterotoma.

Energy allocation is determined by resource availability and trade-offs among traits, and so organisms have to give some traits priority over others to maximize their fitness according to their environment. In this study, we investigated the geographic variations in life history traits and potential trade-offs in populations of the parasitoid Leptopilina heterotoma (Hymenoptera: Figitidae) originating from the north and the south of the Rhône-Saône valley (over a gradient of 300 km, South-East France). We measured a set of traits related to reproduction, maintenance, and mobility using several estimators of each of these main functions determined at different times. We did not find any clear differences between populations from contrasting areas, whereas the southern populations, which were all assumed to be exposed to similar environmental conditions, displayed contrasting patterns of energy allocation. Thus, the most likely explanation seems to be that the evolution of the life history of L. heterotoma is probably shaped by local selective pressures, such as microclimate, microhabitats, or intensity of competition, rather than by regional ecological conditions. Using our study as an example, we discuss the interest of considering several traits and using different ways of measuring them, concluding that multiple measurements should be performed in future studies to ensure the robustness of the results.

Effects of deltamethrin on the specific discrimination of sex pheromones in two sympatric Trichogramma species.

The large amounts of insecticides used for crop protection lead to widespread environmental pollution. Determination of the potential impacts induced by this contamination on key species involved in the equilibrium of ecosystems is therefore a necessity. In this study, we tested the effects of a pyrethroid insecticide, deltamethrin, on the capacity of males from two sympatric Trichogramma species to discriminate the sex pheromones emitted by females of their own species (Trichogramma are parasitoids of Lepidopterous). The impact of an acute exposure as could occur at field edges was evaluated using a dose inducing 20% mortality (LD 20). The impact of a low exposure corresponding to diffuse environmental pollution was evaluated by applying an LD 0.1 (a dose inducing no apparent mortality). For T. semblidis, deltamethrin decreased the specific recognition of sexual pheromones at the higher dose (LD 20) but had no effect on this recognition at the lower dose (LD 0.1). However, deltamethrin decreased the saturation of pheromone receptors at both doses. For T. evanescens, deltamethrin increased the recognition of sexual pheromones at both doses, though not during the same period of observation (at the beginning for the LD 20, at the end for the LD 0.1), but it did not decrease the saturation of the pheromone receptors. These differing results were analyzed considering the behavior of the insects, their level of sensitivity to the insecticide and its mode of action. They provide new insights regarding possible consequences of environmental pollution by insecticides on functional biodiversity.

An inherited virus influences the coexistence of parasitoid species through behaviour manipulation.

The potential role of pathogens or parasites in maintaining species coexistence is well documented. However, the impact of vertically transmitted symbionts, that can markedly modify their host's biology, is largely unknown. Some females of the Drosophila parasitoid Leptopilina boulardi are infected with an inherited virus (LbFV). The virus forces females to lay supernumerary eggs in already parasitised hosts, thus allowing its horizontal transmission. Using two independent experimental procedures, we found that LbFV impacts inter-specific competition between L. boulardi and the related L. heterotoma. While L. boulardi rapidly outcompetes L. heterotoma in the absence of the virus, L. heterotoma was able to maintain or even to eliminate L. boulardi in the presence of LbFV. By forcing females to superparasitise, LbFV induced egg wastage in L. boulardi thus explaining its impact on the competition outcome. We conclude that this symbiont whose transmission is L. boulardi-density-dependant may affect the coexistence of Leptopilina species.

Do variable compensatory mechanisms explain the polymorphism of the dependence phenotype in the Asobara tabida-wolbachia association?

Wolbachia are symbiotic intracellular bacteria, which are classified as reproductive parasites. Although generally facultative, Wolbachia is necessary for Asobara tabida (Hymenoptera), because aposymbiotic females do not produce any offspring. Interestingly, the ovarian phenotype of aposymbiotic females is variable: some females do not produce any eggs, whereas others do produce some eggs, but these are aborted. Here, we show that the ovarian phenotype of aposymbiotic females is highly polymorphic within populations, although dependence remains complete in both cases. We also identified some lines in which aposymbiotic females were able to produce a very few viable offspring, further extending the range of variation observed. These results suggest that various factors actively maintain polymorphism. We demonstrated that Wolbachia is necessary to trigger oogenetic processes, but that the ovarian phenotype was determined by the host only. Phenotypic variation was also correlated with the differential expression of genes controlling iron homeostasis and oxidative stress, which are potentially involved in the evolution of dependence. This suggests that variation in the ovarian phenotype could reflect selection for different levels of compensatory mechanisms in response to Wolbachia infection, and that polymorphism is maintained through selection on different antagonist traits influenced by oxidative stress.

Decrease in fecundity induced by interspecific mating between two Trichogramma parasitoid species.

Trichogramma are used for the biological control of numerous pests. For Trichogramma, as for other insects, the specificity of matings is ensured by several barriers that prevent copulation attempts between insects from different species. We have recently shown that insecticides may totally suppress species recognition that occurs from pheromonal communications between two Trichogramma species, a sublethal effect that will increase mating attempts between two different species. In this work, we have assessed the fitness cost of such interspecific matings and demonstrate that they are very costly for females. After an interspecific mating, females can generate only males because fertilized eggs degenerate (Trichogramma are haplo-diploid species; males are issued from unfertilized eggs and females from fertilized eggs). The resulting offspring are reduced in number by more than half, corresponding to the missing progeny from fertilized eggs. After an interspecific mating, the fecundity of females cannot be restored even if females subsequently mate intraspecifically. These results highlight the strong fitness cost of any event that would decrease the specificity of matings in Trichogramma. Because Trichogramma are key species regulating insect populations, these effects must be considered in the context of sustainable development.

Ecology and life history evolution of frugivorous Drosophila parasitoids.

Parasitoids and their hosts are linked by intimate and harmful interactions that make them well suited to analyze fundamental ecological and evolutionary processes with regard to life histories evolution of parasitic association. Drosophila aspects of what parasitoid Hymenoptera have become model organisms to study aspects that cannot be investigated with other associations. These include the genetic bases of fitness traits variations, physiology and genetics of resistance/virulence, and coevolutionary dynamics leading to local adaptation. Recent research on evolutionary ecology of Drosophila parasitoids were performed mainly on species that thrive in fermenting fruits (genera Leptopilina and Asobara). Here, we review information and add original data regarding community ecology of these parasitoids, including species distribution, pattern of abundance and diversity, host range and the nature and intensity of species interactions. Biology and the evolution of life histories in response to habitat heterogeneity and possible local adaptations leading to specialization of these wasps are reported with special emphasis on species living in southern Europe. We expose the diversity and intensity of selective constraints acting on parasitoid life history traits, which vary geographically and highlight the importance of considering both biotic and abiotic factors with their interactions to understand ecological and evolutionary dynamics of host-parasitoid associations.

[Locomotor activity rhythms in two sympatric parasitoid insects: Eupelmus orientalis and Eupelmus vuilleti (Hyménoptera, Eupelmidae)]. / Rythmes d'activité locomotrice chez deux insectes parasitoïdes sympatriques : Eupelmus orientalis et Eupelmus vuilleti (Hyménoptère, Eupelmidae).

With an automatic image analysis device, we studied the temporal distribution of the locomotor activity of E. orientalis and E. vuilleti during 24 h, and over several days to know whether the activity rhythms of these two Eupelmidae play a role in their competitive interactions. The analysis of locomotor activity rhythms of E. orientalis and E. vuilleti shows that the locomotor activity of both species presents daily cyclic variations. These two Eupelmidae have similar activity rhythms. Displacements of these parasitoids essentially take place during the photophase. But the activity of E. vuilleti is earlier, because the individuals of this species start their activity on average 4 to 5 h earlier than those of E. orientalis. E. vuilleti begins its displacements several hours before the onset of lighting, whereas E. orientalis is active only in the presence of the light. This shift of starting activity is thus a factor allowing these concurrent species to minimize their interactions during the cohabitation period in traditional granaries after the harvests of cowpea.

Comparative life histories and ecophysiology of Drosophila melanogaster and D. simulans.

Numerous laboratory investigations have compared Drosophila melanogaster and D. simulans for various life history traits and fitness related ecophysiological parameters. From presently available information, it is however difficult to get a general comparative pattern describing the divergence of their ecological niches and understanding their demographic success. Two environmental factors seem however to have played a major role: temperature and alcoholic resources. From an ecophysiological approach, D. simulans may be described as generally more sensitive to stresses; other results point to this species as more cold adapted than its sibling; in some cases, however, D. simulans may appear as better adapted to a warm environment. When investigated, ecophysiological traits show a lesser geographic variability in D. simulans than in D. melanogaster. Presently available information does not explain the ecological prevalence of D. simulans in many places with a mild temperate or subtropical climate. This is presumably due to the fact that most comparisons have been done at a single, standard temperature of 25 degrees C. Comparative studies should be undertaken, spanning the thermal ranges of the two species, and the phenotypic plasticity of ecophysiological traits should now be considered.